Characteristics of Annelida: Plesiomorphies and Other Features

1. Symmetry and Size

Annelids are all bilaterally symmetrical animals. They range in size from much less than 1 mm in length to more than 3 m. The smallest annelids described to date would appear to be Neotenotrocha (Dorvilleidae, Eunicida) reaching only a few hundred microns in length (Eibye-Jacobsen and Kristensen, 1994). Also within Eunicida may be found the largest of annelids. A preserved specimen of Eunice (Eunicidae) at the Museum of Natural History in Washington D. C. is close to 3 m in length and around 5 cm in diameter in its preserved and contracted condition. Some earthworms, such as the Giant Gippsland Earthworm (Megascolides australis), can reach 1 m in length. The largest leech in the world, Haementeria ghilianii, is found in South America and reaches more than 40 cm in length.

2. Coelom

Nearly all annelids have a fluid-filled cavity between the outer body wall and the gut, and this is referred to as a coelom (Figure 1). The coelom is often used as a storage area for gametes and acts as a hydrostatic skeleton for locomotion. The coelom (formed via schizocoely) is generally organized as a series of compartments divided by intersegmental septa. This compartmentalization means that if the worm is damaged the coelomic contents will only be lost from a few segments, and locomotion can be maintained. In other annelids there may only be a few septa dividing the coelom. Under these circumstances much more coelomic fluid is lost with injury, and the worm may be severely affected. A number of groups, particularly leeches and a number of small polychaetes, have little or no coelomic space.

3. Body wall

Annelids have a body covered by an external cuticle that is never shed
or molted. Epidermal microvilli secrete a network of fibers that are
in part collagenous and also contain scleroprotein. Chaetae are also
cuticular structures, but contain large amounts of chitin. Beneath the
epidermis and its basal lamina lies a layer of circular muscle. The
circular muscle layer forms a nearly continuous sheath around the body,
except in polychaetes with well-developed parapodia. Beneath the circular
muscle layer lie thick longitudinal muscles. In many annelids the longitudinal
muscles are present as four distinct bands (Fig. 2).

4. Parapodia

Parapodia are unjointed segmental extensions of the body wall found in many polychaetes,
though many do lack these features. They are also absent in Clitellata
and Echiura. Parapodia are equipped with musculature derived mainly
from the circular muscle layer and usually carry chaetae. Parapodia
vary in structure but basically can be considered to consist of two
elements; a dorsal notopodium and a ventral neuropodium (Fig. 3). In
addition to bundles of chaetae, noto- and neuropodia can also have a
variety of cirri and gills. They are most elaborate in actively crawling
or swimming forms where they form large fleshy lobes that act as paddles.
Parapodia of burrowing or tubicolous polychaetes can simply be slightly
raised ridges carrying hooked chaetae called uncini.

5. Nervous system

Annelids have a brain or cerebral ganglion that originates and usually resides in the head. The brain varies in structure, with mobile active forms having the most complex brains, and sessile or burrowing forms having simple brains with little differentiation. It is connected to the ventral nerve cord by the circumpharyngeal connectives, which run down each side of the pharynx. The ventral nerve cord, is usually made up of a pair of cords that are bound together and runs the length of the body (Fig. 4). It varies in thickness and dilates into a ganglion in each segment, from which pairs of segmental nerves pass out to the body wall, muscles and gut.

6. Sense organs

There are six major kinds of sensory structures found in annelids. These include palps, antennae, eyes, statocysts, nuchal organs and lateral organs (Fig. 5). Palps and antennae are located on the head of many polychaetes. In some groups they are both sensory while in others the palps are used for feeding. Nuchal organs are ciliated, paired, chemosensory structures, innervated from the posterior part of the brain. They are present in nearly all polychaetes, and Rouse and Fauchald (1997) suggested that they may represent an apomorphy for Polychaeta. This has been challenged by other authors who suggest that nuchal organs may be an apomorphy for Annelida as a whole and have been lost in Clitellata/Oligochaeta (Purschke et al., 2000). Annelids also have a variety of epidermal sensory cells that may be responsive to light or touch (such as lateral organs).

7. Circulation and respiratory structures

A closed circulatory system is present in most polychaetes (Fig. 6),
Echiura and many clitellates. It is reduced or absent in leeches where
it may be replaced by coelomic canals. A limited circulatory system
in which some of the major blood vessels are present but the distal
capillary vessels are missing is found in a number of polychaete groups.
A circulatory system is absent in many small polychaetes.

8. Segmental organs

In most annelids there are usually two fluid systems, the coelom and
the circulatory system, and both (if present) are involved in the excretion
of waste products. To achieve this excretion there must be ducts to
the exterior, and
these are generally referred to as nephridia. Ducts, known as gonoducts
or coelomoducts, are also required for the transfer of gametes that
develop in the coelom to the outside of the body on maturity. The two
different kinds of ducts are often simply referred to as segmental organs
since determining what kind of duct is present is problematic.

References

Eibye-Jacobsen, D., and R. M. Kristensen. 1994. A new genus and species of Dorvilleidae (Annelida, Polychaeta) from Bermuda, with a phylogenetic analysis of Dorvilleidae, Iphitimidae and Dinophilidae. Zoologica Scripta 23:107-131.

Purschke, G., R. Hessling, and W. Westheide. 2000. The phylogenetic position of the Clitellata and the Echiura - on the problematic assessment of absent characters. Journal of Zoological Systematics and Evolutionary Research 38:165-173.

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Characteristics of Annelida: Plesiomorphies and Other Features
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Greg W. Rouse and Fredrik Pleijel.
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